The First Euthemistid Damsel-Dragonfly from the Middle Jurassic of China (Odonata, Epiproctophora, Isophlebioptera)
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A peer-reviewed open-access journal ZooKeys 261: 41–50The (2013) first euthemistid damsel-dragonfly from the Middle Jurassic of China... 41 doi: 10.3897/zookeys.261.4371 RESEARCH articLE www.zookeys.org Launched to accelerate biodiversity research The first euthemistid damsel-dragonfly from the Middle Jurassic of China (Odonata, Epiproctophora, Isophlebioptera) Yongjun Li1,†, André Nel2,‡, Chungkun Shih3,§, Dong Ren3,|, Hong Pang1,¶ 1 State Key Laboratory of Biocontrol and Institute of Entomology / Key Laboratory of Biodiversity Dynamics and Conservation of Guangdong Higher Education Institutes Sun Yat-Sen University, Guangzhou, China 2 CNRS UMR 7205, CP 50, Entomologie, Muséum National d’Histoire Naturelle, 45 rue Buffon, F-75005 Paris, France 3 College of Life Science, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100048 China † urn:lsid:zoobank.org:author:6F775EC6-130B-42D0-AC5D-62641A20E10A ‡ urn:lsid:zoobank.org:author:98DF555A-16A0-4073-871C-E38BB506C676 § urn:lsid:zoobank.org:author:A49AAC84-569A-4C94-92A1-822E14C97B62 | urn:lsid:zoobank.org:author:D507ABBD-6BA6-43C8-A1D5-377409BD3049 ¶ urn:lsid:zoobank.org:author:9C71F61F-31A9-4296-91A6-B3D67168BD3D Corresponding author: Dong Ren ([email protected]); Hong Pang ([email protected]) Academic editor: M. Engel | Received 21 November 2012 | Accepted 8 January 2013 | Published 24 January 2013 urn:lsid:zoobank.org:pub:04114911-498B-4400-9396-E6ED89ABEC4D Citation: Li Y, Nel A, Shih C, Ren D, Pang H (2013) The first euthemistid damsel-dragonfly from the Middle Jurassic of China (Odonata, Epiproctophora, Isophlebioptera). ZooKeys 261: 41–50. doi: 10.3897/zookeys.261.4371 Abstract Sinoeuthemis daohugouensis gen. et sp. n. is the first record of the isophlebiopteran family Euthemistidae from Middle Jurassic of northeast China, while previously this family was restricted to the early Late Jurassic Kazakhstan. This new finding allows us to emend the family diagnosis with hindwing characters. This new species shows a mixture of characters alternatively present in different genera of the two families Euthemistidae and Sphenophlebiidae. Keywords Euthemistidae, Sphenophlebiidae, gen. et sp. n., Middle Jurassic, China, mixture of characters Copyright Yongjun Li et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 42 Yongjun Li et al. / ZooKeys 261: 41–50 (2013) Introduction The Mesozoic Isophlebioptera Bechly, 1996 is a very large clade subdivided into four subgroups: Euthemistidae, Parazygoptera, Selenothemistidae, and Isophlebiida. The family Euthemistidae Pritykina, 1968 comprises one genus and two species (both based on forewings). Our knowledge on the Euthemistidae remains indigent and its exact position is somewhat uncertain, even if it is probably very inclusive in the Iso- phlebioptera. Here we describe a new genus and a new species collected from the Jiulongshan Formation, Middle Jurassic of Daohugou, Inner Mongolia, China, where abundant well-preserved fossil insects are found, including 19 reported orders so far, and some of the oberservations are of great importance (e.g. Ren et al. 2009, 2010a, 2010b, 2010c; Gao et al. 2012; Gu et al. 2012; Shi et al. 2011). The new genus is closely related to the genus Euthemis Pritykina, 1968 and for the first time we show the hindwing characters of the family Euthemistidae. The new fossil has a mixture of characters alternatively present in different genera of the two families Euthemistidae and Sphenophlebiidae. Therefore, this new finding is of great importance for future clarification of the relationships between the Euthemistidae, Sphenophlebiidae, and other Parazygoptera. Furthermore, the new specimen expands the age and distribution of Euthemistidae from early Late Jurassic Karatau (Kazakhstan) to Middle Jurassic northeast China. Material and method The study is based on one specimen (No. CNU-ODO-NN2012004, positive imprint and negative imprint) housed in the Key Laboratory of Insect Evolution and Envi- ronmental Changes, Capital Normal University, Beijing, China. The specimen was examined with a Leica MZ12.5 dissecting microscope and illustrated with the aid of a drawing tube attached to the microscope. Line drawings were made using Adobe Photoshop CS graphic software. We use the following standard abbreviations: AA, anal vein; AP, anal posterior; Ax0 Ax1 Ax2, primary antenodal cross-veins; CuAa, distal branch of cubitus anterior; CuAb, proximal branch of cubitus anterior; IR1, IR2, intercalary radial veins; MAa, distal branch of median anterior; MAb, posterior branch of median anterior; MP, me- dian posterior; N, nodus; O, oblique vein; Pt, pterostigma; RA, radius anterior; and RP, radius posterior; ScP, subcosta posterior. We follow the taxonomy of Isophlebiida indicated by the phylogenetic system of Bechly (1996), but do not accept all the syna- pomorphies he proposed (see discussion below). The first euthemistid damsel-dragonfly from the Middle Jurassic of China... 43 Systematic palaeontology Order Odonata Fabricius, 1793 Clade Isophlebioptera Bechly, 1996 Superfamily Isophlebioidea Handlirsch, 1906 Family Euthemistidae Pritykina, 1968 http://species-id.net/wiki/Euthemistidae Genus included. type genus Euthemis Pritykina, 1968 and Sinoeuthemis gen. n. Emended familial diagnosis. Several long intercalary veins between IR1 and RP1, and between IR1 and RP2, as well as between RP3/4 and IR2, and between IR2 and RP2 (intercalaries parallel to main longitudinal veins without apparent origin on them, but originating in cross-venation); extremely narrow postdiscoidal area (in fore- wings and probably also in hindwings); not petiolate; numerous secondary antenodal crossveins between anterior wing margin and ScP distal of Ax2; discoidal cells opened in forewing and closed in hindwing; hindwing subdiscoidal area transverse, posteriorly closed, short and broad, with vein CuAb makes a strong angle with AA; hindwing gaff (basal CuA before its branching) not very long; RP2 aligned with subnodus; crossveins in hindwing postdiscoidal space are not very long and not oblique. Genus Sinoeuthemis gen. n. urn:lsid:zoobank.org:act:1DE29A69-7318-4D84-B0EE-209593108C8A http://species-id.net/wiki/Sinoeuthemis Diagnosis. Wings relatively short and very short CuA with weak posterior branches in both fore and hindwings. Etymology. Named after Sinica, Latin name for China and Euthemis, the type genus. Gender feminine. Sinoeuthemis daohugouensis sp. n. urn:lsid:zoobank.org:act:6783C7B8-8791-48B0-9722-010427CECC6E http://species-id.net/wiki/Sinoeuthemis_daohugouensis Figure 1 Material. Holotype specimen No. CNU-ODO-NN2012004. Diagnosis. As for the genus. Description. A body with a thorax, abdomen, head, two legs and forewings and hindwing articulated. Body (Fig. 1; Fig. 2D,F) 53.0 mm long (from head to anal appendages); head 5.1 mm long, 5.2 mm wide, with broad eyes, 1.7 mm long, well separated, 1.0 mm apart in the mid level; thorax about 8 mm long, max width 6.5 mm; 44 Yongjun Li et al. / ZooKeys 261: 41–50 (2013) abdomen about 3.8 mm wide in the mid part, slightly narrowed at the end; cercus and epiproct very short; there is no secondary genital structure on segment 2 and anal area rounded (female). Forewing (based on negitive imprint, two forewing fragments combined; Fig. 2A,B), preserved with basal half, 22.4 mm long; no petiole (AA and AP separate at wing base); one row of cells between posterior wing margin and AA; AA parallel to MP + Cu; median and submedian areas free; a curved strong vein CuP be- tween submedian and subdiscoidal areas, in a distal position just basal of arculus; sub- discoidal space free of cross-veins, transverse; discoidal space basally opened; RP+MA nearly straight, separated at nearly a right angle from RA in arculus; distance between base of RP and point of separation between MAa and MAb 0.4 mm, RP and MA well parallel; MAb 0.9 mm long, well aligned with distal free part of CuA; CuA separates from MP 4.3 mm from wing base and directed towards posterior wing margin for 0.6 mm; distal free part of CuA strong, CuA distally fused with AA; CuA divided into a very short CuAb directed towards posterior wing margin and CuAa basally more or less parallel to posterior wing margin and distally delimitating a short and narrow cubi- to-anal area, with 1-2 posterior branches and 1-2 rows of cells at its broadest part; apex of CuA slightly distal level of base of RP3/4; area between CuA and MP with one row of cells; distal of apex of CuA, area between MP and posterior wing margin very long and broad; MP nearly straight, certainly reaching posterior wing margin well distal of nodus level; MAa more or less parallel with MP, nearly straight in its preserved part; postdiscoidal area with one row of cells, 1.0 mm wide near discoidal cell and narrow- ing distally; Ax0 not preserved; Ax1 0.6 mm basal of arculus, disposed obliquely to ScP and R + MA, Ax2 2.4 mm distal of arculus, with inverted obliquity; eight preserved secondary antenodal cross-veins between C and ScP distal of Ax2; 13 visible secondary antenodal cross-veins between ScP and RA distal of Ax2; 15 preserved cross-veins in area between RA and RP between arculus and subnodus; base of RP3/4 5.0 mm distal of arculus, closer to arculus than to nodus; base of IR2 close to that of RP3/4, 3.8 mm distally; no visible antefurcal cross-vein in space